Persistence of Apache Trout Following Wildfires in the White Mountains of Arizona

Home , Apache trout, Mount Baldy (Arizona)

PERSISTENCE OF APACHE TROUT FOLLOWING WILDFIRES IN THE WHITE MOUNTAINS OF ARIZONA

Jonathan Long

Wildfires are a natural disturbance that can Mountain suggest that this landscape has rejuvenate and rebuild trout habitat by stim- experienced stand-replacing wildfires, ulating flood scour and deposition (Gress- because extensive stands of aspen (Populus well 1999; Bisson et al. 2003). However, tremuloides) often reflect past wildfires in several case studies from Arizona and New mixed-conifer forests (Swetnam et al. 2001; Mexico have shown severe reductions in fish Margolis 2003). However, a recent study of populations shortly after wildfires (Neary et stand-replacing high-elevation fires (Mar- al. 2005). The consequences of wildfire are a golis 2003) did not include the White Moun- particular concern in the mountainous wat- tains, in part because other areas offered ersheds that harbor Apache trout (Oncorhyn- clearer historical evidence of such fires (Ellis chus gilae ssp. apache) and Gila trout (Onco- Margolis, personal communication). rhynchus gilae ssp. gilae). Populations of these Recent climatic conditions and historical federally protected species are scattered management practices have increased the among small stream reaches that are isolated likelihood of large and severe wildfires in by the presence of non-native trouts, inter- the White Mountains. Widespread insect mittent flows, thermal regimes, natural infestations, abetted by warm winters and barriers, and barriers to exclude non-native dry summers, have killed huge swaths of trout (Brown et al. 2001; Rieman et al. 2003; spruce in bands around Mount Baldy USFWS 2006). Wildfires have extirpated (Lynch 2004). The combination of increased seven populations of Gila trout since 1989, forest fuels and warming temperatures over impeding recovery efforts (Propst et al. 1992; the past century has increased the likelihood USFWS 2006). Emergency evacuation of of wildfires across vegetation zones in Ari- populations threatened by fire has become zona mountains (Cocke et al. 2005). Re- an important element of Gila trout recovery searchers have asserted that past timber (Anderson 1992; USFWS 2006), and has been harvest and road construction activities have recommended as a “fundamental manage- rendered habitats more vulnerable to post- ment approach” for conserving isolated pop- fire impacts; this combination of threats has ulations of rare native fishes (Rinne 2004). motivated many land managers and com- However, because evacuation is costly and munity members to express concern that stressful to fish, managers need to know large wildfires in the White Mountains when it is necessary to prevent population could jeopardize valued resources such as losses. Apache trout (Abrams 2005). Debates over The extent of the threat to Apache trout how to manage forests in the face of such from wildfire has not been clear, since large risks need to be guided by analyses of wild- wildfires have not struck its ancestral home fire effects within local landscape contexts on Mount Baldy in the White Mountains of (Rieman et al. 2003). Arizona in several decades (Dieterich 1983; Two streams (Grant Creek and KP Creek) Gomez and Tiller 1990). Place names such as whose high-elevation watersheds have Aspen Butte, Aspen Ridge, and Burnt recently experienced wildfires (the Steeple 220 Jonathan Long fire of July 2003 and KP fire of May 2004) are (Schaffner and Reed 2005). Cannon et al. especially important because they represent (2003) found that slopes greater than 30 per- ancestral habitat for native trout (Figure 1). cent and soils with increased permeability Native trout had been collected from KP and organic matter content were associated Creek in 1904, although the taxonomic with increased probability of fire-induced identity is uncertain because the original debris flows. Debris flows are also strongly specimens have been lost (Miller 1950). KP related to the occurrence of post-fire storm Creek was later stocked with a strain of events (Cannon 2001), but it is difficult to Apache trout from an unconnected stream predict such events. The availability of light- as part of recovery efforts (Dowling and ly burned refugia and absence of barriers are Childs 1992). Researchers have concluded two factors that help fish avoid impacts that the present-day trout populations in from fires (Burton 2001; Bisson et al. 2003). both KP Creek and Grant Creek are hybrid- Previous research in the White Moun- ized with rainbow trout (Figure 2; Rinne tains indicates that landscape attributes such 1985; Dowling and Childs 1992; USFWS as topography, geology, and vegetation type 2003). The current recovery plan is to stock need to be considered when evaluating fish these streams with the Spruce Creek lineage habitat (Long and Medina 2005). Within a of Gila trout, which appears to be an inter- region, mean basin elevation and slope help mediate form between Gila and Apache to explain variation in post-fire flooding trout (Riddle et al. 1998; USFWS 2003). The (Schaffner and Reid 2005). Therefore, com- impact of wildfire on the two creeks was paring landscape attributes of the study thus not a major concern from a conserva- streams to those designated for Apache tion standpoint, but their response to wild- trout recovery would help in evaluating fires could yield insights into how wildfires whether their post-fire responses would be might affect other streams that have been representative. Extrapolating the responses designated for recovery of Apache trout. of two streams to a much larger and more The severity of a burn within a stream’s diverse region is speculative, but it could watershed is a primary factor in predicting help managers who must act on limited the impact of fires on aquatic systems. information concerning the risk of wildfires Burton (2001) found that 60–88 percent of to Apache trout. the watersheds of streams that experienced high levels of post-fire debris flooding had METHODS burned at high severity; only 14–37 percent This study examined the upper reaches of of the watersheds of streams that did not KP and Grant Creeks, perennial tributaries experience post-fire debris flooding were of the Blue River (Figure 1). A five-person burned at high severity. Cannon (2001) has team from the Rocky Mountain Research reported that the percent of watershed area Station (RMRS) sampled fish populations burned at moderate to high severity is an and habitat conditions at seven 50 m long important predictor of debris flows, and sampling sites in KP and Grant Creeks in Schaffner and Reed (2005) successfully pre- June of 2004 (Figure 3), after the KP fire was dicted increases in peak flows by calculating contained. Fish populations were resampled area burned at moderate to high severity. at six of the sites one year later. The team Managers can easily evaluate burn severity attempted to relocate sites that had been in the aftermath of a wildfire, making it an sampled for fish in September of 1995 by the important and accessible tool for predicting Arizona Game and Fish Department fish persistence in different settings. (AGFD) as part of their General Aquatic A variety of factors besides burn charac- Wildlife Surveys (GAWS) program. The teristics affect the responses of stream eco- team successfully located three of those systems to wildfires, including human monitoring sites (labeled 22, 20, and 19 from influences and natural landscape attributes upstream to downstream) in the upper Jonathan Long 221

Figure 1. Locations of Apache trout streams and the study streams in the White Mountains of east- central Arizona. reaches of Grant Creek, although AGFD had with nets to prevent fish from escaping sampled fish only at the middle site. In KP during sampling and each was sampled Creek, the original site markers could not be three times using the depletion method located, so the team approximated the three (Zippin 1958). Lengths of all fishes were uppermost sites (24, 23, and 22 from up- recorded and used to calculate mean lengths stream to downstream) based on the with 95 percent confidence intervals. The descriptions of channel gradient and intra- maximum likelihood population estimate station distances recorded on the 1995 and 95 percent confidence interval were survey forms. Fish populations were sam- calculated at each sampling site using the pled using backpack-mounted electro- Microfish 3.0 computer program (Van shocking gear. Each reach was blocked off Deventer and Platts 1989). 222 Jonathan Long

Figure 2. Hybridized Apache trout collected in KP Creek, June 2004.

Land Resource Information System served Landscape Attributes of Study Sites to characterize site geology (Figure 4). The Watersheds of the study sites were deline- two major geologic types are felsic volcanic ated using the hydrologic modeling tools in formations along the upper slopes of Mount ArcMap version 9 (ESRI, Redlands, CA). Baldy and mafic volcanic formations along Burn severity information (based on differ- the lower slopes and adjacent plateaus. enced normalized burn ratio data calculated Slopes were calculated using ArcMap to from satellite images) provided by the process 10 m digital elevation models of the Apache-Sitgreaves National Forest was region (Figure 5). Data for the vegetation incorporated into the geographic database, layer were provided by the U.S. Geological and the data were used to calculate the per- Survey’s GAP analysis program. In the three cent of each watershed burned at moderate major mixed-conifer forest types that pre- to high severity. dominate within the range of Apache trout Mapping landscape attributes of KP and (Figure 6), the dominant tree species are Grant Creek served to compare them to the Engelmann spruce (Picea engelmannii), Doug- 36 streams that the Apache Trout Recovery las fir (Pseudotsuga menziesii), and ponderosa Team has identified as priorities for conser- pine (Pinus ponderosa). vation of pure strains of Apache trout within Maps and elevations in the draft recovery its ancestral range (Ruiz and Novy 2000). A plan and Arizona Game and Fish Depart- coarse-scale map of Arizona (Reynolds 1988) ment’s GAWS reports indicated the upper available in digital format from the Arizona and lower limits of habitat in each stream, Jonathan Long 223

Figure 3. Map of fire impact study sites with overlay of burn severity for the KP and Steeple fires.

and in the absence of upstream limits, the pacts if only the lower part of the watershed beginning of a solid blue line on topographic were burned. maps approximated the start of habitat. The blue-line method provides only a rough RESULTS approximation of suitable habitat (Svec et al. KP Creek 2005), but it facilitates comparison of the The estimates of trout population at the general landscape characteristics of Apache three reaches sampled in KP Creek in 2005 trout streams. Figure 7 classifies the streams were higher than or equal to the population into three landscape types based on the counts at their approximate locations when characteristics at the upper end of habitat surveyed 10 years earlier (Table 1). The nu- within a stream. The highest-elevation habi- meric data from 2004 were misplaced and tat could serve as a refuge from wildfire im- not recovered, but the memories of the 224 Jonathan Long

Figure 4. Geology of the White Mountains, revealing that Apache trout streams originate from the felsic slopes of Mount Baldy (light area) or on adjacent mafic plateaus (dark gray and medium gray areas with triangular stippling).

surveyors were that the numbers and sizes of fish were similar to those collected in Grant Creek 2005. Photographs taken during sampling in In Grant Creek, temporal comparisons were 2004 indicated that fishes were present in limited by the fact that fish had been sam- sizes ranging from at least 95 to 195 mm. pled only at one site (20) prior to the fire. The percent of severely and moderately That sample was collected in September of burned watershed above each site increased 1995. Resampling was conducted in early from 4.7 percent at the upstream site to 8.4 June in both 2004 and 2005. The population percent at the middle site and to 19.1 percent estimate within reach 20 in 1995 was twice at the downstream site. The team observed the levels estimated in 2004 and 2005 (Table no evidence of recent debris flows or chan- 1). However, the fish in the 1995 sample nel incision at the sites in 2004 or 2005, but a were only half the length of those in 2004 very large debris fan was observed in the and 2005. The apparent abundance of small channel < 100 m downstream from the low- fish could reflect seasonal variation, since ermost site in 2004 (Figure 8). It emanated the late summer sampling could have from a drainage that burned at high severity collected young-of-year fish. Sampling at a in the previous year’s Steeple fire, which downstream site (19) yielded higher popu- had burned into the KP watershed. lation estimates and smaller sizes compared Jonathan Long 225

Figure 5. Topography of the White Mountains with darker shading indicating steeper slopes.

to site 20 in both 2004 and 2005 (Table 1). A fish movements upstream. A debris fan reach between sites 19 and 20 (19.5) that was below site 19 also appeared to have formed sampled only in 2004 had sizes and numbers another potential barrier. Site 22 had the of trout similar to site 20 (Table 1). Sampling highest burn severity and was the only sam- at the uppermost site (22) indicated that no pled reach where the channel was incised. fish were present even though that reach had flowing water in 2004 and 2005. Landscape Comparison The percent of watershed burned at mod- KP and Grant Creeks are similar to most erate to high severity at the sampling sites Apache trout streams in that they flow was higher than at sites in KP Creek (Table through mixed-conifer forests in mafic 1). The channel along Grant Creek was canyons. However, their upper watersheds impacted by debris flows emanating from (where the study sites were located) are side canyons in several locations. These relatively steeper than most Apache trout flows formed alluvial fans that confined the watersheds (Figure 5). Habitat in 11 Apache channel and induced upstream aggradation. trout streams extends into the high-elevation Water flowed underneath piles of woody (over 3000 m) spruce zone on Mount Baldy, debris in the reach above site 20 (Figure 9). and seven others harbor Apache trout only Although the reach itself was not heavily within the lower elevation (below 2500 m) impacted, the pile had the potential to block ponderosa pine zone (Figure 7). Table 1. Trout population estimates, lengths, and burn severity at sites on Grant Creek and KP Creek, White Mountains, Arizona (x = no data, NA = not applicable).

Population Estimate (Total Catch to Upper 95% CI) Mean Length ± 95% CI (in mm) % Watershed Burned at 9/95 6/04 6/05 9/95 6/04 6/05 Moderate (pre- (before runoff (1 year (pre- (before runoff (1 year to High Site fire) from fire) after fire) fire) from fire) after fire) Severity

Grant 22 x 0 0 NA NA NA 29.8% Grant 20 28 (28-29) 14 (14-15) 15 (15-15) 84 ± 11 174 ± 11 174 ± 28 23.2% Grant 19.5 x 13 (13-14) x x 179 ± 7 x 22.4% Grant 19 x 20 (18-27) 23 (19-35) x 158 ± 11 154 ± 21 28.3% KP 24 4* x 11 (11-13) x x 115 ± 32 4.7% KP 23 12* x 12 (12-13) x x 129 ± 17 8.4% KP 22 25* x 33 (33-35) x x 140 ± 12 19.1%

*1995 data for KP creek were reported only as total fish caught, not a maximum likelihood estimate based upon three-pass depletion. Jonathan Long 227

Figure 6. Vegetation types in the White Mountains, with darker shades representing high-elevation communities with naturally less frequent fire regimes.

Figure 7. Bar chart representing the approximate elevation ranges of trout habitats in study streams arranged by geologic types and shaded by approximate vegetation zone. The whiskers above each bar denote the elevation at the summit of each watershed. 228 Jonathan Long

Figure 8. Large debris flow in KP Creek below site 22, emanating from a small area that was severely burned by the Steeple fire the year previously.

flows were not observed in the year after the DISCUSSION less severe KP fire. These results are consis- The differences in fish populations and tent with the hypothesis that the extent of lengths in Grant Creek before and after the moderate to high burn severity is an impor- fires could reflect seasonal variation (i.e., tant predictor of physical impacts such as fewer young fish in the June samples) or debris flows. Since trout did survive in both post-fire impacts that might have inhibited streams, the results indicate that wildfires recruitment of young fish. The data from that are severe enough to induce debris both creeks are insufficient to evaluate flows do not necessarily extirpate trout whether the fire caused changes in abun- populations. dance. They do demonstrate that fish per- Table 2 compiles previous research by the sisted in both KP and Grant Creeks for at Rocky Mountain Research Station and other least the first year of post-fire runoff, which published reports of wildfire impacts on fish Neary et al. (2005) have suggested is a criti- populations within watersheds in eastern cal timeframe for extirpations to occur. The Arizona. The percent of each watershed Steeple fire appeared to have induced debris burned at moderate or high severity was flows along both KP and Grant Creeks, calculated using the methods described pre- resulting in debris piles that could have viously. The KP fire also burned the head- created barriers to fish movement. Such waters of Raspberry Creek, which flows Table 2. Impacts of fires on fish habitat in east-central Arizona based on various case studies between 1990 and 2004.

Watershed Dominant Area Burned Hydro- Effects on Location Streams Upland at Moderate– Geomorphic Fish Fire (in Arizona) Examined Vegetation High Severity Impacts Populations References

Dude (1990) Mogollon Rim Bonita Creek Pine > 75%* Debris flows and Extirpation Rinne 1996 channel incision Ellison Creek Pine > 75%* Debris flows and Extirpation channel incision Dude Creek Pine > 50%* Debris flows and Extirpation channel incision (1 trout found 1 year later)

White Springs Mogollon Rim White Spring Pine ~ 75% Debris flows and Extirpation Long & Endfield 2000 (1996) channel incision

Steeple (2003) White Mtns Grant Creek Douglas fir/ 28.3% Debris flows and Trout persisted This study (site 19) pine channel incision Raspberry Creek Pine 7.0% — Trout persisted

Rodeo-Chediski Mogollon Rim Limestone Pine 72% Debris flows and Extirpation Long & Burnette 2004 (2002) Canyon channel incision

Steeple (2003) White Mtns Grant Creek Douglas fir/ 30% Debris flows and Trout persisted This study (site 19) pine channel incision Raspberry Creek Pine 7.0% — Trout persisted USFWS 2006

KP (2004) White Mtns KP Creek Douglas fir/ 19.1% Not observed Trout persisted This study (site 22) pine 1 year post-fire

Three Forks White Mtns N. Fork E. Fork Douglas fir/pine, 3.5% Short-term turbidity Trout persisted Unpub. data, J. Rinne, (2004) Black River mixed grass/shrub increase due to ash C. Carter & J. Long Boneyard Creek 4.8% (RMRS) W. Fork E. Fork 6.4% of Black River

*Estimated based on fire perimeter maps and observations by J. Leonard, an RMRS technician who has studied the Dude fire. 230 Jonathan Long

Figure 9. Debris flow in Grant Creek above site 20.

down a steep, south-facing slope through fire and afterwards. The post-fire samples in ponderosa pine forest (Figures 6 and 7). In 2004 and 2005 showed that fish persisted response, federal and state agencies evacu- and no changes in channel morphology ated a portion of the Gila trout population were observed (unpublished data of J. that had been introduced to the stream, and Rinne, C. Carter, and J. Long on file at the the U.S. Forest Service set a backfire to inhib- RMRS in Flagstaff). it the spread of the fire. A maximum of only The Steeple, KP, and Three Forks fires 7 percent of the watershed burned at moder- had more moderate impacts compared to ate to high severity; the U.S. Fish and Wild- earlier wildfires on ponderosa pine forests life Service reported that trout survived in along the Mogollon Rim west of the White the creek (USFWS 2006). Mountains. Several of the earlier fires, in- The Three Forks fire burned 3200 hectares cluding the Dude fire of 1990 (Rinne 1996), in the relatively flat headwaters of the East the White Springs fire of 1996 (Long and Fork of the Black River (Figures 2 and 6). Endfield 2000), and the Rodeo-Chediski fire Only 4–6 percent of each tributary drainage of 2002 (Long and Burnette 2004; Long et al. burned at moderate to high severity (Table 2005), induced fish extirpations and major 2). Historical data from AGFD’s GAWS sur- impacts to stream channels including chan- veys indicate that brown trout and various nel avulsion and rapid incision. In each case, warmwater species such as desert sucker localized extirpations of fish populations and speckled dace were present in samples occurred within the first couple of years at from all periods, although Apache trout sites where more than 50 percent of the were not. The samples were highly variable watershed area burned at moderate to high between sampling periods both before the severity (Table 2). Jonathan Long 231

The Aspen fire of 2003 burned most of drought, historical fire suppression, and the upper Sabino Creek watershed and thus heavy insect damage fueled the 11,800 ha triggered an evacuation of rare Gila chubs Nuttall Complex fire in spruce-fir and from Sabino Creek (Tobin 2004). Post-fire mixed-conifer forest on Mount Graham in debris flows in the canyon substantially 2004 (Koprowski et al. 2005). The fire, which modified aquatic habitats and initially burned 59 percent of the watershed of Mari- appeared to eliminate all aquatic vertebrates jilda Creek at moderate to high severity, in the canyon (USDA 2004). However, re- induced extreme flooding and debris flows searchers subsequently discovered several (Schaffner and Reed 2005), demonstrating adult and numerous young Gila chubs in a that high-elevation streams in east-central large pool (Tobin 2004). Forty-seven percent Arizona are vulnerable to high-severity of the watershed above that reach burned at wildfire given the current climate and stand moderate to high severity. conditions. However, management options These case studies of fire impacts are con- may be limited because fuel reduction treat- sistent with a hypothesis that the proportion ments are unlikely to mitigate fire hazards of moderate to high severity burn is a useful in this naturally dense forest type (Schoen- predictor of impacts to fishes and their nagel et al. 2004). habitats. Specifically, extirpations have been The spruce zone on Mount Baldy, which reported for streams where more than half harbors many of the larger populations of of the watershed burned at moderate to high Apache trout, was the source of three of the severity. By consistently collecting pre- and remaining pure lineages of this fish (Ord, post-fire data, future studies could test such East Fork White River, and Smith). Water- a threshold of burn severity. sheds in this zone have slopes in excess of 30 Despite its potential utility, an index of percent, relatively coarse-textured geologic burn severity is a simple metric that ignores formations, and soils with high organic- many factors that are likely to influence matter content (Long et al. 2003). If the rela- post-fire stream impacts. However, land- tionships reported by Cannon et al. (2003) scape classifications can help to control for apply to this region, then post-wildfire important factors such as watershed steep- debris flows could be more likely to occur in ness, soil erodibility, and availability of streams on Mount Baldy than in streams on refugia. Managers need to tailor their man- the adjacent basaltic plateaus. On the other agement strategies to the historic fire regime hand, several of the high-elevation streams, within particular forest types (Schoennagel including Ord Creek, East Fork of the White et al. 2004). Accordingly, the typology used River, and Big Bonito Creek, have multiple in Figure 7 may be helpful in planning man- perennial tributaries that could provide agement of Apache trout habitats. refuge for trout.

Spruce Zone Mixed Conifer Zone Stand-replacing wildfires in the subalpine (Douglas fir Dominated) zone dominated by spruce are normally The KP and Steeple fires resulted in relative- infrequent due to high moisture (Veblen et ly large, mixed-severity burns within the al. 1991; Swetnam and Baisan 1996), but Douglas fir zone following several years of stand-replacing fires have been reported in drought conditions; mixed-severity fire spruce-fir communities in northern New regimes are typical in this zone (Rollins et al. Mexico during severe droughts in the mid 2000; Margolis 2003). Dieterich (1983) found eighteenth century (Margolis 2003). The risk that the mean fire return interval in this for- of stand-replacing wildfires in this zone may est type in the White Mountains is 22 years, be increasing due to the combination of and stand-replacing fires appear to be un- drought, warming, and insect outbreaks, common. However, severe droughts com- which has increased fuel loads in the form of bined with heavy fuel loading and insect needles and dead trees. A combination of outbreaks could trigger more severe fires in 232 Jonathan Long this forest type. Two such fires occurred in CONCLUSIONS 1951—the Escudilla fire east of Mount Baldy KP and Grant Creeks are similar in geology, (Savage and Mast 2005) and the McKnight topography, and vegetation to the majority fire in New Mexico (Rollins et al. 2000). The of streams planned for recovery of Apache McKnight fire left a legacy of incised and trout. These streams extend into mixed coni- unstable channels prone to scouring (Me- fer forests where mixed-severity wildfires dina and Martin 1988), and harsh floods such as the KP and Steeple fires are typical. have repeatedly diminished the populations Fish extirpations have been reported from of Gila trout that were introduced into streams in drier, lower-elevation forest types McKnight Creek after the fire (USFWS 2003). where wildfires have been more severe, but That outcome suggests that wildfires in this long-term fire history studies suggest that zone could have more severe consequences high-severity wildfires do occur in high- than were observed during this study. elevation forest types during extended dry periods. Trout populations persisted follow- Ponderosa Pine Zone ing the mixed-severity wildfires in KP and Fire-scar studies have suggested that Grant Creeks, indicating that evacuation of wildfires in relatively dry ponderosa pine populations may be unnecessary when a forests along the Mogollon Rim of Arizona watershed is not severely burned. While and the Gila Mountains of New Mexico have many factors can influence the likelihood of historically been primarily low ground fires fish persistence, burn severity can be deter- with return intervals of less than 10 years mined rapidly using satellite imagery. Con- (Swetnam and Baisan 1996). More than two- sequently, until more confirmatory studies thirds of Apache trout streams extend into are conducted, that metric may help mana- the ponderosa pine zone. Meanwhile human gers evaluate whether to evacuate Apache alteration of fire regimes in those forest trout populations that are threatened by types may have increased the occurrence of wildfires. high-severity crown wildfires (Brown et al. 2001; Savage and Mast 2005), which caused ACKNOWLEDGMENTS the fish extirpations reported from the Mo- The U.S. Forest Service provided funding for gollon Rim. Not all wildfires in ponderosa this research through the National Fire Plan. pine forests in Arizona and New Mexico W. Wall with the Apache-Sitgreaves Nation- have caused such severe impacts, though; al Forest and M. Lopez and K. Meyer with for example, Earl and Blinn (2004) reported the Arizona Game and Fish Department that recent wildfires in ponderosa pine shared historical data on fish and fires in the forest in New Mexico generated relatively region. I give special thanks to C. Carter, J. short-lived impacts to streams. The risks of Leonard, S. Kelly, and J. Rinne for their high-severity wildfire are lower when such assistance in collecting the field data for this stands are kept less dense (Rollins et al. study. I also thank three anonymous review- 2000). Nevertheless, as is the case in other ers whose detailed comments substantially forest types, severe fires may also have improved the manuscript. occurred naturally during warmer, drier periods (Pierce et al. 2004). In the relatively LITERATURE CITED moist ponderosa pine forests found at higher elevations, mixed-severity fire re- Abrams, J. 2005. Report on a needs assessment for collaborative landscape planning in the White gimes naturally predominate and treatments Mountains of Arizona. Ecological Restoration to reduce the risk of high-severity wildfire Institute, Northern Arizona University. Avail- are less likely to be effective (Schoennagel et able online at http://www.forestera.nau.edu/ docs/wm_needs_assessment_final.pdf. al. 2004). Jonathan Long 233

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